MOSQUITO ECOLOGY Field Sampling Methods. Second Edition
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1 MOSQUITO ECOLOGY Field Sampling Methods Second Edition
2 MOSQUITO ECOLOGY Field Sampling Methods SECOND EDITION M. W. SERVICE Vector Biology and Control, Liverpool School of Tropical Medicine, Liverpool, UK SPRINGER-SCIENCE+BUSINESS MEDIA. BV.
3 First edition published by Elsevier Science Publishers Ltd 1976 Second edition published by Elsevier Science Publishers Ltd 1993 Reprinted by Springer-Science+Business Media, BV Springer Science+Business Media Dordrecht Originally published by Chapman & Hall in 1993 Softcover reprint of the hardcover 2nd edition ISBN ISBN (ebook) DOI / Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permission in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page. The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made. A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data available
4 Preface to Second Edition It is relatively easy to produce a first edition, all that one needs is self-discipline, an inexhaustible supply of paper, a good library and an understanding spouse. A second edition requires something more. You have to read through the entire first edition and in so doing begin to realise that probably no one else has ever done so. You then begin to be overwhelmed by the amount of new information that you have managed to discover on your subject and ponder on just how selective you should be in integrating it into the new edition. At this point you may reflect that Dr Samuel Johnson said 'A man may turn over half a library to make one book'. After which you may try and console yourself with another of his quotations, namely, 'What is written without effort is in general read without pleasure'. I have generally adopted the layout of the previous edition but have expanded sections that, although not directly concerned with sampling, are nevertheless associated with it. For example, I have given more details on oviposition attractants, larval growth retardant factors, and aggregation of populations, and now include summaries of methods to identify blood-meals, and accounts of vectorial capacity and the calculation of inoculation rates. I have added a small final chapter, which one of my students described as containing 'junk data'. But after having written all other chapters I was left with information on recent books on mosquito ecology and behaviour that contain information relevant to sampling, accounts of attempts to model mosquito populations, and publications containing important mathematical and statistical considerations, that did not easily fit into anyone chapter, but were applicable to many. So Chapter 12 was created for this type of data. In the 17 years since the First Edition was published there has been a proliferation of publications on mosquito ecology, especially on subjects such as estimating adult survivorship and on mosquitoes colonising tree-holes and other natural container-type habitats. During the intervening years many mosquito workers have broadened their outlook and adopted or modified ecological principles and techniques used by ecologists for their own studies on mosquitoes. At the same time mosquitoes have attracted the attention of 'pure ecologists' who have brought new ideas to the study of mosquito ecology, although admittedly they have usually concentrated on species breeding in discrete container habitats. Such tractable microhabitats make it easier to sample and study mosquitoes v
5 VI PREFACE TO SECOND EDITION living in them. Several of these papers are highly mathematical and are frightening to those who have no love for mathematics. I have tried to understand such papers and present the basics so that entomologists can apply the procedures and analyse their data without needing to read the original papers; which for some may be difficult to obtain. It has proved a daunting task reading the enormous volume of literature for this Second Edition, which contains some 1800 additional references. Only a few of the older ones could be deleted. Finally, it is a pleasure to thank my wife, Wendy, for typing and retyping the manuscript, checking the references, making the indices, proof-reading, and encouraging me to continue when I felt I was losing the battle with such a wealth of new information. M. W. Service Liverpool December 1991
6 Preface to First Edition This book has been written for the field worker, whether he is mainly concerned with mosquito control and wishes, for example, to learn more about the use of light-traps for monitoring pest mosquitoes or maybe improve his larval sampling procedures, or whether he is more interested in ecology and population dynamics of mosquitoes. Descriptions of traps and procedures, and methods of analysing results, have been described mostly in sufficient detail to enable those with limited access to the original publications to understand and use the methods. It has proved impractical to describe all the many different types of traps and methods that have been used to collect mosquitoes. I have tried therefore to present a selection of the more useful and interesting techniques while at the same time introducing others that, although little used in mosquito studies, appear to have potential. In describing apparatus and techniques I have retained the units of measurement given in the original descriptions. Although this results in some descriptions being in metric and others in non-metric units it avoids the 'odd' amounts which result from converting original measurements, frequently chosen for convenience, to the metric scale. M. W. Service Liverpool Vll
7 Contents Preface to Second Edition Preface to First Edition. Acknowledgements Useful Conversion Data v vii xi xiii Chapter 1 SAMPLING THE EGG POPULATION 1 Chapter 2 SAMPLING THE LARVAL POPULATION 75 Chapter 3 SAMPLING THE ADULT RESTING POPULATION 210 Chapter 4 SAMPLING ADULTS WITH NON-ATTRACTANT TRAPS. 291 Chapter 5 SAMPLING ADULTS BY ANIMAL BAIT CATCHES AND BY ANIMAL-BAITED TRAPS 349 Chapter 6 SAMPLING ADULTS WITH CARBON DIOXIDE TRAPS, LIGHT-TRAPS, VISUAL ATTRACTION TRAPS AND SOUND TRAPS 499 Chapter 7 SAMPLING THE EMERGING ADULT POPULATION 611 Chapter 8 EXPERIMENTAL HUT TECHNIQUES FOR EVALUATING INSECTICIDES 637 Chapter 9 MARK-RECAPTURE TECHNIQUES AND ADULT DISPERSAL. 652 ix
8 x CONTENTS Chapter 10 ESTIMATION OF THE MORTALITIES OF THE IMMATURE STAGES AND ADULTS 752 Chapter 11 INDICES OF ASSOCIATION BETWEEN SPECIES AND SPECIES DIVERSITY. 890 Chapter 12 ECOLOGICAL AND STATISTICAL PUBLICATIONS. 914 Author Index 925 Mosquito Species Index 963 Subject Index 971
9 Acknowledgements Grateful acknowledgements are given to the authors and publishers of figures, tables and formulae that I have used in original or modified form. In all instances references have been given to the authors in the text, tables or figures. I am indebted to the World Health Organization and the Office de la Recherche Scientifique et Technique Outre-Mer for permission to refer to their unpublished mimeographed reports and documents. I would like to thank the editors of the following for permission to reproduce redrawn figures, modify published tables and in a few instances make short quotations: American Journal of Epidemiology, American Journal of Tropical Medicine and Hygiene, American Midland Naturalist, Annals of the Entomological Society of America, Annals of Tropical Medicine and Parasitology, Biometrika, Bulletin of Entomological Research, Bulletin of the Pan-American Health Organization, Bulletin of the Society for Vector Ecology, Cahiers ORSTOM, Serie Entomologie Medicale et Parasitologie, Canadian Entomologist, Canadian Journal of Zoology, East African Virus Institute Report, Ecology, Ecological Entomology, Endemic Disease Bulletin of Nagasaki University, Entomologia Experimentalis and Applicata, Entomological News, Entomologist's Gazette, Entomologist's Monthly Magazine, Environmental Entomology, Folia Parasitologia, Heredity, Hydrobiologia, Japanese Journal of Sanitary Zoology, Journal of Animal Ecology, Journal of Applied Ecology, Journal of the Australian Entomological Society, Journal of Communicable Diseases, Journal of Economic Entomology, Journal of the Entomological Society of Southern Africa, Journal of Hygiene, Journal of Medical Entomology, Journal of Parasitology, Journal of Tropical Medicine and Hygiene, Medecine Tropicale, Medical and Veterinary Entomology, Mosquito News, Nature, Pacific Entomologist, Physiological Entomology, Proceedings of thelinnean Society of London, Oecologia, Proceedings of the California Mosquito and Vector Control Association, Proceedings of the South London Entomological and Natural History Society, Researches on Population Ecology, Revista de Saude Publica, Transactions of the Royal Entomological Society of London, The Entomologist, Tropical Biomedicine, Tropical Medicine, and to Methuen & Co, W. B. Saunders & Co., the Entomological Society of America, Defence Research Board of Canada and the Commonwealth Agricultural Bureaux. I am greatly indebted to the following, arranged in alphabetical order, who xi
10 xii ACKNOWLEDGEMENTS kindly supplied me with diverse information; J. C. Beier, L. R. Boobar, W. E. Bradshaw, A. N. Clements, S. E. Cope, F. M. Eskafi, R. P. Fall, M. G. Fletcher, J. E. Freier, H. Ginsberg, A. K. Githeko, Y. H. Heng, J. W. Hock, F. R. Holbrook, G. J. Hunt, T. Ikeshoji, T. Ishii, P. G. Jupp, B. H. Kay, D. L. Kline, M. Labuda, S. V. Landry, K. J. Linthicum, Lu Bao Lin, S. E. Lunt, J. McNelly, C. L. Meek, M. V. Meisch, R. P. Meyer, M. M. Milby, C. Mitchell, M. Mogi, L. E. Munstermann, J. K. Nayar, M. L. Niebylski, J. K. Olson, W. C. Reeves, W. K. Reisen, P. Reiter, M. Renshaw, J. A. Rozendaal, B. L. Sharp, J. A. Shemanchuk, G. R. Shidrawi, W. F. Snow, B. Tesh, E. D. Walker, A. Warburg, A. A. Weathersbee, H. H. Yap, M. G. Yates and Zairi bin Jaal. I would also like to thank Natalija V. Nikolaeva for translating Russian papers and discussing with me the work of several Russian entomologists. In addition the following most kindly read and criticised various sections, M. Begon, M. H. Birley, W. L. Bidlingmayer, C. Dye, J. R. Fry, M. T. Gillies, M. J. Lehane, P. Mason, A. Saul and L. R. Taylor. Finally, it is a pleasure to thank Kenneth Mellanby whose idea it was that I should write the First Edition of this book.
11 Useful Conversion Data To convert Inches to centimetres Centimetres to inches Yards to metres Metres to yards Miles to kilometres Kilometres to miles Square inches to square centimetres Square centimetres to square inches Square yards to square metres Square metres to square yards Acres to hectares Hectares to acres Ounces to grams Grams to ounces Pounds to kilograms Kilograms to pounds Fluid ounces to millilitres Millilitres to fluid ounces Pints to litres Litres to pints Gallons to litres Litres to gallons Cubic feet to cubic metres Cubic metres to cubic feet Cubic yards to cubic metres Cubic metres to cubic yards Imperial gallons to US gallons US gallons to Imperial gallons Multiply by xiii
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